Placodermi
Placodermi (from the Greek πλάξ = plate and δέρμα = skin, literally "plate-skinned") is a class of armoured prehistoric fish, known from fossils, which lived from the Silurian to the end of the Devonian period. Their head and thorax were covered by articulated armoured plates and the rest of the body was scaled or naked, depending on the species. Placoderms were among the first jawed fish; their jaws likely evolved from the first of their gill arches. Placoderms are paraphyletic, and consist of several distinct outgroups or sister taxa to all living jawed vertebrates, which originated among their ranks. This is illustrated by a 419-million-year-old fossil, Entelognathus, from China, which is the only known placoderm with a type of bony jaw like that found in modern bony fishes. This includes a dentary bone, which is found in humans and other tetrapods. The jaws in other placoderms were simplified and consisted of a single bone. Placoderms were also the first fish to develop pelvic fins, the precursor to hindlimbs in tetrapods, as well as true teeth. Paraphyletic groupings are problematic, as one can not talk precisely about their phylogenic relationships, their characteristic traits and literal extinction. 380-million-year-old fossils of three other genera, Incisoscutum, Materpiscis and Austroptyctodus, represent the oldest known examples of live birth. In contrast, one 2016 analysis concluded that placodermi are likely monophyletic. The first identifiable placoderms appear in the fossil record during the late Llandovery epoch of the early Silurian. The various groups of placoderms were diverse and abundant during the Devonian, but became extinct at the end-Devonian Hangenberg event 358.9 million years ago. Many placoderms, particularly the Rhenanida, Petalichthyida, Phyllolepida, and Antiarchi, were bottom-dwellers. In particular, the antiarchs, with their highly modified, jointed bony pectoral fins, were highly successful inhabitants of Middle-Late Devonian freshwater and shallow marine habitats, with the Middle to Late Devonian genus, Bothriolepis, known from over 100 valid species. The vast majority of placoderms were predators, many of which lived at or near the substrate. Many, primarily the Arthrodira, were active, nektonic predators that dwelled in the middle to upper portions of the water column. A study of the arthrodire Compagopiscis published in 2012 concluded that placoderms (at least this particular genus) likely possessed true teeth contrary to some early studies. The teeth had well defined pulp cavities and were made of both bone and dentine. However, the tooth and jaw development were not as closely integrated as in modern gnathostomes. These teeth were likely homologous to the teeth of other gnathostomes. One of the largest known arthrodires, Dunkleosteus terrelli , was 6 m (20 ft) long, and is presumed to have had a large distribution, as its remains have been found in Europe, North America and possibly Morocco. Some paleontologists regard it as the world's first vertebrate "superpredator", preying upon other predators. Other, smaller arthrodires, such as Fallacosteus and Rolfosteus, both of the Gogo Formation of Western Australia, had streamlined, bullet-shaped head armor, strongly supporting the idea that many, if not most, arthrodires were active swimmers, rather than passive ambush-hunters whose armor practically anchored them to the sea floor. Some placoderms were herbivorous, such as the Middle to Late Devonian arthrodire Holonema, and some were planktivores, such as the gigantic, 8 m (26 ft) long arthrodire, Titanichthys. It was thought for a time that placoderms became extinct due to competition from the first bony fish and early sharks, given a combination of the supposed inherent superiority of bony fish and the presumed sluggishness of placoderms. With more accurate summaries of prehistoric organisms, it is now thought that they systematically died out as marine and freshwater ecologies suffered from the environmental catastrophes of the Devonian/Carboniferous extinction events. Fossil record The earliest identifiable placoderm fossils are from China and date to the early Silurian. At that time, they were already differentiated into antiarchs and arthrodires, as well as other, more primitive, groups. Earlier fossils of basal Placodermi have not yet been discovered. The Silurian fossil record of the placoderms is both literally and figuratively fragmented. Until the discovery of Silurolepis (and then, the discoveries of Entelognathus and Qilinyu), Silurian-aged placoderm specimens consisted of fragments. Some of them have been tentatively identified as antiarch or arthrodire due to histological similarities; and many of them have not yet been formally described or even named. The most commonly cited example of a Silurian placoderm, Wangolepis of Silurian China and possibly Vietnam, is known only from a few fragments that currently defy attempts to place them in any of the recognized placoderm orders. So far, only three officially described Silurian placoderms are known from more than scraps: • the basal antiarch Silurolepis, from the Ludlow epoch of Yunnan, China, known from an almost complete thoracic armor. Currently, Placodermi are divided into eight recognized orders. There are two further controversial orders: One is the monotypic Stensioellida, containing the enigmatic Stensioella; the other is the equally enigmatic Pseudopetalichthyida. These orders are considered to be basal or primitive groups within Placodermi, though their precise placement within the class remains unsure. Fossils of both are currently known only from the Hunsruck lagerstatten. Category:Palaeozoic monsters Category:Fish